mele n. sp., a new species of freshwater goby from Vanuatu and New Caledonia (Teleostei, , Sicydiinae), and comments about amphidromy and regional dispersion

Philippe KEITH Muséum national d’Histoire naturelle, Département Milieux et Peuplements aquatiques, Ichtyologie, case postale 26, 57 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected]

Gérard MARQUET 96 rue de Richelieu, F-75002 Paris (France)

Marc POUILLY Antenne IRD, UR 131, Universidad Mayor de San Simón, Unidad de Limnología y Recursos Acuáticos, Casilla postale 2352, Cochabamba (Bolivia)

Keith P., Marquet G. & Pouilly M. 2009. — Stiphodon mele n. sp., a new species of freshwater goby from Vanuatu and New Caledonia (Teleostei, Gobiidae, Sicydiinae), and comments about amphidromy and regional dispersion. Zoosystema 31 (3) : 471-483.

ABSTRACT Stiphodon mele n. sp. is described on the basis of material collected from Vanuatu and New Caledonia. It is distinguished from all other congeners in having 9 segmented rays in the second dorsal fi n, usually 13 pectoral rays, 36-41 fi ne tricuspid premaxillary teeth, 2 small symphyseal teeth in female vs. 2-4 stout teeth in males, predorsal scales absent, and low number of scales in transverse back (3-7) and lateral series (14-21). Th is new species spawns in freshwater, the free embryos drift downstream to the sea where they undergo a planktonic phase, KEY WORDS before returning to the rivers to grow and reproduce: it is an amphidromous Teleostei, species. As many other Sicydiinae, Stiphodon mele n. sp., is endemic to Vanuatu Gobiidae, Stiphodon mele n. sp., and New Caledonia. Numerous factors occur to control the dispersion of the Vanuatu, amphidromous species. It is necessary to consider all of these factors to explain New Caledonia, the existence of many endemic species and the broad or small distribution of freshwater, amphidromy, the species in the Indo-Pacifi c area. Th e management and the conservation of new species. Stiphodon mele n. sp. have to be necessarily regional.

ZOOSYSTEMA • 2009 • 31 (3) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 471 Keith P. et al.

RÉSUMÉ Stiphodon mele n. sp. une espèce nouvelle de gobie d’eau douce du Vanuatu et de Nouvelle-Calédonie (Teleostei, Gobiidae, Sicydiinae) et commentaires sur l’amphidromie et la dispersion régionale. Stiphodon mele n. sp. est décrit à partir de matériel récolté au Vanuatu et en Nouvelle-Calédonie. Il se distingue des autres espèces du genre par la présence de 9 rayons segmentés dans la seconde nageoire dorsale, 13 rayons aux nageoires pectorales, 36 à 41 dents prémaxillaires tricuspides, 2 petites dents symphyséales chez les femelles et 2 à 4 chez les mâles, aucune écaille dans la région prédorsale et moins d’écailles en série transverse postérieure (3-7) et en série latérale (14-21). Cette espèce nouvelle pond en eau douce, les embryons libres sont entraînés vers la mer où ils vont mener une vie planctonique avant de revenir vers les MOTS CLÉS rivières pour les coloniser, grossir et s’y reproduire : c’est une espèce amphidrome. Teleostei, Comme beaucoup d’autres Sicydiinae, Stiphodon mele n. sp., est endémique Gobiidae, Stiphodon mele n. sp., du Vanuatu et de la Nouvelle-Calédonie. De nombreux facteurs contrôlent Vanuatu, la dispersion des espèces amphidromes. Il est nécessaire de considérer tout ces Nouvelle-Calédonie, facteurs dans la région Indo-Pacifi que pour expliquer l’existence des endémiques eau douce, amphidromie, et leur répartition large ou réduite. La gestion et la conservation de Stiphodon nouvelle espèce. mele n. sp. doivent être nécessairement régionales.

INTRODUCTION dertaken on the major hydrosystems of Grande-Terre and Loyalty Islands (Marquet et al. 2003). Th ese Th e freshwater ichthyofauna of Vanuatu is both rich inventories were completed between 2004 and 2008 and varied but remained poorly sampled until 1998 by further specifi c studies, particularly in Panié and, as a result, little has been published on these Mountain, Forgotten Coast and Belep Islands. over the centuries (see Keith et al. 2004a). In Th ese recent surveys in Vanuatu and New Cale- 1998, the Environment Unit of Vanuatu sponsored donia have permitted to add new records and to a survey of freshwater environments throughout the discover new species (Watson et al. 2001, 2002, islands of Vanuatu (Gerbeaux et al. 1998; Keith et 2005; Keith et al. 2004a, b, 2007a; Lord & Keith al. 2002, 2004a, b, 2007a, b). Between 2002 and 2008). Th e purpose of this paper is to provide a 2008, the Muséum national d’Histoire naturelle, description of Stiphodon mele n. sp., a freshwater Paris (MNHN) conducted numerous surveys of fi shes goby known from Vanuatu and New Caledonia occurring in freshwater streams and rivers on the islands and to discuss about its larval dispersion. Th e genus of Maewo, Santo, Tanna, Ambae, Efate, Malekula and Stiphodon Weber, 1895 has three haemal spines Gaua, and in 2006, an extensive eff ort was made in prior to the fi rst anal pterygiophore, a narrow based Santo rivers with the SANTO 2006 Expedition (for pelvic disk and pelvic fi n adherent to belly between a narrative of the expedition, see Bouchet et al. 2008, fi fth rays only, the tongue fused to the fl oor of the and for a review of the geography and natural history mouth and the dorsal tip of ascending process on of Santo, we refer to Bouchet et al. in press). premaxilla narrower than the process below. Th e freshwater ichthyofauna of New Caledonia is also both rich and varied but remained poorly known until the end of the 20th century. Between MATERIAL AND METHODS 1998 and 2003, the MNHN conducted, with the provincial and territorial authorities, an exhaustive Methods follow those in Watson et al. (2005) and inventory of freshwater fi shes. Inventories were un- Keith et al. (2007a). All lengths of specimens are

472 ZOOSYSTEMA • 2009 • 31 (3) A new species of goby (Teleostei, Gobiidae) from Vanuatu and New Caledonia

expressed with a dial caliper in standard length (MNHN 2008-1921) — Santo, Patunar’s Doline, (SL) to the nearest tenth of a millimetre (mm). Jaw 14.IX.2006, Pouilly coll., 2 ♂♂, 1 ♀ 21.1-25.3 mm SL length is measured from anterior tip of upper jaw (MNHN 2008-1922). — Santo, Patunar resurgence, 15.IX.2006, Pouilly coll., 1 ♀ 23.8 mm SL (MNHN to posterior edge of maxilla. Scales in a lateral series 2008-1923). are counted from upper pectoral base and along the New Caledonia. North Province, Panie tributary, Keith middle of the body laterally to the central hypural et al. coll., 1 ♂ 26.2 mm SL (MNHN 2008-1924). base. Body depth is measured from anterior base Remarks: seven specimens were collected from Vanuatu of second dorsal fi n to belly, this measurement is and New Caledonia totaling fi ve males and two females with a size range of 21.1-27.2 mm SL, largest male taken only from males as females vary considerably 27.2 mm SL. All specimens caught were adults and not from gravid to non gravid state. Transverse series juveniles. Th e metamorphosis was fi nished, the males back, refers to scales counted from the fi rst scale were mature (urogenital papilla well diff erentiated and anterior to second dorsal fi n, in a diagonal manner, colourful specimens parading when caught) and the posteriorly and ventrally to the anal fi n base or female gravid. ventralmost scale. Transverse series forward, refers COMPARATIVE MATERIAL. — Th e new species is compared to scales counted from the fi rst scale anterior to in text and tables with species of nearby areas in particular second dorsal fi n, in a diagonal manner, anteriorly Stiphodon rubromaculatus Keith & Marquet, 2007, and ventrally to the centre of belly or ventralmost S. kalfatak Keith, Marquet & Watson, 2007, S. birdsong Watson, 1996, S. hydoreibatus Watson, 1999, S. sur- scale. Zigzag series, refers to scales on the narrowest rufus Watson & Kottelat, 1995, S. tuivi Watson, 1995, region of the caudal peduncle counted from the S. astilbos Ryan, 1982 and S. sapphirinus Watson, Keith & dorsalmost scale to the ventralmost scale in a zigzag Marquet, 2005 because these have 13-14 pectoral rays (alternating) manner. and 9 segmented rays in the second dorsal fi n. Abbreviations used to represent the cephalic sensory Material for S. rubromaculatus, S. birdsong, S. surrufus, S. hydoreibatus, S. tuivi and S. sapphirinus is that listed pore system follow (1986). Th e material cited in Keith & Marquet (2007), and for S. kalfatak and in the present paper is deposited in the Muséum S. astilbos in Keith et al. (2007a). national d’Histoire naturelle, Paris (MNHN). Diagrammatic illustrations of the head and the ETYMOLOGY. — Th e species is named mele to honour Mele waterfall in Efate where the fi rst specimen was urogenital papilla are not provided in this work be- found (noun in apposition). cause there is no great morphological variation among the species so far described from streams fl owing DIAGNOSIS. — A combination of characters distinguishes into the Pacifi c Ocean (Watson et al. 2005). Stiphodon mele n. sp. Th e species has 13 pectoral rays, 9 segmented rays in the second dorsal fi n and 36-41 Counts and morphometrics are summarized in premaxillary teeth. Th ere are no scales in predorsal the Appendix. midline. Th e most anterior scale along midline is below anterior part of second dorsal fi n. Males have more and larger symphyseal teeth than females (2-4 vs. 2). Th e SYSTEMATICS species has 3-7 scales in transverse backward series. Th e typical colouration of males is orange; the female is greyish to brownish. Family GOBIIDAE Linnaeus, 1758 Subfamily SICYDIINAE Gill, 1860 Genus Stiphodon Weber, 1895 DESCRIPTION Number of premaxillary teeth, scale counts, and Stiphodon mele n. sp. morphometrics in Stiphodon mele n. sp. and related (Fig. 1) species are given in Appendix (Tables 1; 2-5 and 6-15 respectively). Below, the holotype counts are TYPE MATERIAL. — Holotype: Vanuatu. Efate, Mele given fi rst followed in brackets, if diff erent, by the waterfall, 22.VII.2002, Keith and Keith coll., ♂ 27.2 mm SL (MNHN 2008-1920). paratypes counts. Paratypes: Vanuatu. Gaua, Solomul river, 21.VII.2005, Dorsal fi ns VI-I,9, fi rst dorsal fi n separate from Keith, Marquet and Keith coll., 1 ♂ 21.1 mm SL and of same height as second dorsal fi n; spines not

ZOOSYSTEMA • 2009 • 31 (3) 473 Keith P. et al.

elongate or fi lamentous. Anal fi n I,10 and directly few brownish pigment. Background of head orange, opposite to second dorsal fi n. Pectoral fi n counts 13 greyish or yellowish; occipital region with brownish rays, uppermost rays extending beyond membrane pigment; preoperculum behind eye with a small but not appearing feathery or silky, lowermost 1 yellowish patch. First dorsal fi n spines with 2 to 5 or 2 rays simple; fi n oblong with posterior margin blackish dots evenly spaced; second dorsal fi n spines rounded. Caudal fi n with 13 (13-14) branched and rays with 2 or 3 blackish dots evenly spaced. rays, posterior margin rounded. Pelvic disc always Caudal fi n with slightly dusky brown crescent-shaped with 1 spine and 5 stout and heavily branched margin posteriorly; anal fi n with a thin clear distal segmented rays. Fifth rays joined together in their margin, below distal margin a band of brownish entire length forming a strong adhesive disc; disc pigment; pelvic disk with few brownish pigment adherent to belly between fi fth rays only; between along distal margin. spines a strong fl eshy frenum. Scales in lateral series 14 (14-21), those on caudal Females. Background of body yellowish; light peduncle are ctenoid and they become cycloid be- dusky black band extending midlaterally from low the second dorsal fi n. Most anterior scale along posterior to pectoral base ending as a spot posterior midline below anterior part of second dorsal fi n. to hypural base. Another light dusky brown band Scales in transverse backward series 3 (3-7). Scales extends from posterior to eye ending at upper caudal in transverse forward series 0 (0-7) with scales well rays; belly and abdomen brownish. Background developed at origin of second dorsal fi n when present. of head greyish; blackish brown band from snout Scales in zigzag series 6 (5-6). No scales in predorsal and upper lip to central pectoral base. First dorsal midline in males (0-1 in female). Females have a fi n clear except for fi ne brownish pigment along few small cycloid scales close to urogenital papilla anterior and posterior edges of each spine; second and anus. Usually no scales on belly of males. Head, dorsal fi n clear, spine and rays with dusky brown breast and pectoral base are without scales. pigment; caudal fi n clear, rays without pigment; Premaxillary teeth 36-41, fi ne and tricuspid, tri- anal fi n mostly clear; pelvic disk free of pigment; dent I-shaped with central cusp longer than lateral pectoral fi n clear. cusps. Dentary symphyseal teeth in males 3 (2-4) (females, 2), conical to canine I-shaped, stronger Colour in life and larger. Males. Two diff erent patterns. First one (Fig. 1), Cephalic sensory pore system always A, B, C, generally bright orange on side of head and body. D, F, H, K, L, N and O; pore D is singular, all Cheeks and nose orange or yellow. Caudal fi n others are paired. Oculoscapular canal separated hyaline with a reddish part at its base. Th e second into anterior and posterior canals between pores pattern is more greyish on entire body. Posterior H and K. Cutaneous sensory papillae developed margin of scales blackish. All fi ns hyalines, some over lateral and dorsal surfaces of head. with dark dots: 2 to 5 black dots on each rays of Sexual dimorphism well developed with adult fi rst dorsal fi n; 2 or 3 black dots on each rays of males always having bright orange or greyish col- second dorsal fi n. oration, presence of dots on dorsal fi ns and longer second dorsal and anal fi ns than female. Urogeni- Females. Body generally hyaline or greyish; dusky tal papilla in males somewhat rectangular with a markings as in preservation. Belly whitish to yel- rounded distal tip, while in female rectangular with lowish. tiny distal fi mbriate projections. COMPARISON Colour in preservation Stiphodon mele n. sp. diff ers from S. rubromaculatus Males. Background of body pale orange or yellowish; and S. surrufus because it has more premaxillary many scales along and above midline below second teeth (36-41 vs. 27-32/25-36), lower scale counts dorsal fi n with ctenii generally blackish; belly with in transverse back series (3-7 vs. 7-9/10-14) and in

474 ZOOSYSTEMA • 2009 • 31 (3) A new species of goby (Teleostei, Gobiidae) from Vanuatu and New Caledonia

FIG. 1. — Stiphodon mele n. sp., holotype ♂ 27.2 mm SL (MNHN 2008-1920), Efate, Mele waterfall, 22.VII.2002, Vanuatu, Keith and Keith coll. Photo: É. Vigneux. transverse forward series in males (0-1 vs. 1-4/0-12). DISTRIBUTION It diff ers from S. sapphirinus, S. tuivi, S. hydoreibatus, Known from rivers of Gaua, Efate and Santo in S. kalfatak and S. astilbos, because it has lower scale Vanuatu, Mont Panié tributaries and Bélep Islands counts (14-21 in lateral scales vs. 21-35/23-39/23- in New Caledonia. 35/30-34/24-29; 3-7 in transverse back series vs. 10-12/10-14/9-10/8-9/9; 0-7 in transverse forward ECOLOGY series vs. 7-15/12-19/10-14/7-8/12-16). Stiphodon Like other Sicydiinae, Stiphodon mele n. sp. is found mele n. sp. diff ers from S. birdsong in having less in clear, high gradient streams with rocky bottom. It scales in transverse back series (3-7 vs. 8-15), more lives mainly on the riverbed, on top of rocks but it premaxillary teeth (36-41 vs. generally 28-37) is also seen swimming in open water in the current and a shorter caudal peduncle in length (10-16 between rocks or in large pools. Th is species was vs. 17-23). observed from 10 to 100 m high in altitude.

KEY TO VANUATUAN AND NEW CALEDONIAN SPECIES OF STIPHODON WEBER, 1895 1. 13 or 14 pectoral rays. Size < 3.5 cm ...... 2 — 15 pectoral rays. Size > 3.5 cm ...... Stiphodon atratus 2. Second dorsal fi ns with 10 branched rays ...... Stiphodon rutilaureus — Second dorsal fi ns with 9 branched rays ...... 3 3. First and second dorsal fi ns with longitudinal black stripes ...... Stiphodon mele n. sp. — First and second dorsal fi ns without longitudinal black stripes ...... 4 4. First dorsal fi n of male non fi lamentous; less than 9 scales (or 9 scales) in transverse back series ...... 5 — First dorsal fi n of male fi lamentous; more than 9 scales in transverse back series ...... Stiphodon sapphirinus 5. Less than 35 premaxillary teeth; 12-16 scales in transverse forward series ... Stiphodon astilbos — More than 35 premaxillary teeth; 7 or 8 scales in transverse forward series ...... Stiphodon kalfatak

ZOOSYSTEMA • 2009 • 31 (3) 475 Keith P. et al.

COMMENTS ABOUT AMPHIDROMY (Keith et al. 2005b); Parasicydium is restricted to AND REGIONAL DISPERSION West Africa (Pezold et al. 2006) and Akihito, a re- cently described genus (Watson et al. 2007; Keith In the Indo-Pacifi c region, insular river systems are et al. 2007b) seems to be restricted to the West of colonised by Gobiidae, in particular Sicydiinae, Pacifi c Ocean. with a life cycle adapted to the conditions in these Within these genera we observe diff erent distri- distinctive habitats, which are young oligotrophic bution patterns (Keith et al. 2009). Among them, rivers subject to extreme climatic and hydrological we can clearly distinguish a specifi c one sharing seasonal variation. Th ese species spawn in freshwater, endemic species between New Caledonia and Van- the free embryos drift downstream to the sea where uatu. Th e discovery of a new species of Stiphodon they undergo a planktonic phase, before return- endemic to this region is very interesting from both ing to the rivers to grow and reproduce (Keith biogeographical and conservation points of view. 2003; Keith et al. 2008), hence they are qualifi ed We assume that there are dispersion and exchange as amphidromous (McDowall 2007). Th e practical of larvae between New Caledonia and Vanuatu, details of their biological cycle and the parameters owing to their amphidromous life cycle, depending leading to such extreme evolution in amphidromous on the species and the duration of the larval stage. gobies are poorly known, despite the fact that these Indeed, the only possibility for the dispersion of gobies contribute most to the diversity of fi sh the species and the colonisation of islands is the communities in the Indo-Pacifi c insular systems, marine planktonic larval phase. Th e duration of and have the highest levels of endemism (Keith et this planctonic stage varies from 91 to 265 days al. 2006; Lord & Keith 2008). Th e amphidromous for several species of Sicydiinae of the Pacifi c and life cycle is, for these species, the major way for Caribbean region (Keith 2003; Murphy & Cowan dispersal and it allows them to colonise new rivers 2007). Th e strength and direction of marine cur- and new islands. But amphidromy also highlights rents as well as the duration of the planktonic phase the fact that there are factors which can limit or could infl uence the dispersion ability, the species favour the dispersion of species and enabling them distribution area, and the evolution of the group. to be either “endemic” or “cosmopolit”. In a recent study Keith et al. (2005a) have shown Among amphidromous gobies, the Sicydiinae that an amphidromous species ( lago- Bleeker, 1874 subfamily comprises eight genera cephalus (Pallas, 1767)), present in New Caledonia (Stiphodon [30 species]; Sicyopus Gill, 1863 [15]; and Vanuatu, has a broad distribution and occurs Günther, 1861 [12]; Guichenot, over a range of 18 000 km in the Indian and Pa- 1864 [2]; Sicyopterus Gill, 1860 [30]; cifi c oceans. Th e overall pattern of distribution and Valenciennes, 1837 [16]; Akihito Watson, Keith & phylogenetic relationship suggest that the lineages Marquet, 2007 [2] and Parasicydium Risch, 1980 leading to endemic species in this genus originated [1]), and is the most diversifi ed of the group with earlier than S. lagocephalus. Th e latter seems to be a nearly 110 species distributed in Indo-Pacifi c, West secondary migrant species, having gradually colo- Africa, Central America and Caribbean areas (Keith nised both Indian and Pacifi c Oceans. 2003). All these genera have specifi c distribution and But the duration of the larval phase and the refl ect a specifi c evolution in the areas concerned. strength of the marine current are not the only fac- Indeed, Sicydium is distributed throughout Carib- tors involved. Th e substrate specifi city could play a bean, Central America and West Africa (Pezold et major role. As part of Gondwana, New Caledonia has al. 2006), Sicyopterus is distributed in Indo-Pacifi c some specifi c substrates not found in volcanic areas area from West Indian Ocean to East Pacifi c one like Vanuatu. So, there are more specifi c endemic (Keith et al. 2005a), Stiphodon, Sicyopus, Lentipes amphidromous fi shes of New Caledonia (Marquet are distributed from East Indian Ocean to East Pa- et al. 2003) living on ultramafi c substrate (perido- cifi c (Watson et al. 2001, 2002; Keith & Marquet tite) like many insect groups and reptiles (Keith et 2007), Cotylopus is restricted to West Indian Ocean al. 2009). But there are also other amphidromous

476 ZOOSYSTEMA • 2009 • 31 (3) A new species of goby (Teleostei, Gobiidae) from Vanuatu and New Caledonia

gobies living in short, quick and oxygenated rivers marine currents (Radtke et al. 2001). Th e survival of the Mont Panié in the northeast, on metamorphic of the species depends also on the ability of existing substrate, that are endemic to New Caledonia and populations to provide enough larvae to maintain Vanuatu (Keith et al. 2009). appropriate numbers of adults. Finally, numerous factors occur to control the On islands, the impact of humans on aquatic dispersion of the species and the specifi city of each habitats is highly signifi cant, particularly on estuarine island. Planes & Galzin (1997) divide factors aff ect- habitats which are crucial to amphidromous species. ing genetic structure and larval dispersion of coral Th ese have to undertake two migrations between reef populations in the Pacifi c Ocean into three freshwater and the sea. Th e success of such a life major groups: 1) functional factors which are related cycle – i.e. production of larvae for downstream to the ecology and the biology of each species (this migration after hatching and restocking rivers includes parameters such as reproductive behaviour with post-larval and juvenile upstream colonisa- or duration of the pelagic larval phase); 2) physical tion after recruitment in freshwaters – depends on factors including the eff ect of oceanic currents and/ maintaining the mountain-ocean corridor open to or the geomorphological structure of the biotope; allow movements between both habitats (Radtke and 3) historical factors such as colonisation or et al. 2001). Moreover, anthropogenic impacts to extinction linked to sea level variation, which will streams, pollution, or deforestation could diminish modify directly the population structure. the fl ow rates and hereby the substrate type, shift- In the case of amphidromous species, it is neces- ing from hard rocky substratum to silt, preventing sary to consider all of these factors to explain the Sicyopterus species to stick on the substrate and to existence of many endemic species and the broad or migrate upstream (Lord & Keith 2006, 2008). A small distribution of the species in the Indo-Pacifi c change in fl ow rates can also alter the algal type of area. It is indeed probable that a complex system the riverbed, directly infl uencing the survival of including marine currents, duration of the larval Sicydiin species. Man-made developments on these phase, paleohistory of the area (period of emergence streams can alter larval dispersion and therefore of the islands, vicariousness and variation of the sea the recruitment success (Keith et al. 2008). But if level, etc.), in addition to the particularities of the managers rehabilitate degraded rivers, these could be islands (altitude, substratum, age, velocity of river, recolonised, as the amphidromous life cycle allows etc.), are at the origin of the structure observed the dispersion at sea and the recolonisation. (Keith et al. 2009). Finally, seasonal variables (e.g., rainfall, drought, Th e mechanism of larval production, retention, fl oods, typhoons) could have also a major impact dispersal, and recruitment to freshwater are both on the survival of populations: biological events governed by biological and physiological proc- such as reproduction, spawning, and the dispersal esses (Murphy & Cowan 2007). Th e current state of larvae, are dependent on these events and are of knowledge on the life cycles of amphidromous synchronised with them (Keith 2003). gobies (biology, ecology), the length of the larval Just discovered, this new species seems to be phase and the part it plays in the dispersal of larvae, already very rare. It is therefore necessary to un- is of direct relevance for management and conser- derstand its biology to the best of our ability, and vation. Th is management and the conservation of to develop regional management and restoration amphidromous species is necessarily regional, due strategies between New Caledonia and Vanuatu in to dispersion mode, and must take into account order to preserve amphidromous gobies (Keith & both the dependency of adult populations on the Marion 2002). larval pool for replacement, and the contribution of each reproductive population to the larval pool. Th e length of the marine phase might increase or Acknowledgements reduce the probability of fi nding a river for colo- A part of this work was fi nancially supported by nisation, as will the strength and the direction of the New Caledonian Government, the North

ZOOSYSTEMA • 2009 • 31 (3) 477 Keith P. et al.

Province, the Société française d’Ichtyologie, the of amphidromous Gobiidae of the Indo-Pacifi c and French Ministery of Ecology and the MNHN. Th e the Caribbean regions. Journal of Fish Biology 63: SANTO 2006 Expedition was organized by Muséum 831-847. KEITH P. & M ARION L. 2002. — Methodology for national d’Histoire naturelle, Paris, Pro Natura drawing up a Red List of threatened freshwater fi sh International (PNI), and Institut de Recherche in France. Aquatic Conservation 12: 169-179. pour le Développement (IRD). It operated under KEITH P. & M ARQUET G. 2007. — Stiphodon rubromacu- a permit granted to Philippe Bouchet (MNHN) latus, a new species of freshwater goby from Futuna by the Environment Unit of the Government of island (Teleostei: Gobioidei: Sicydiinae). Cybium 31 (1): 45-49. Vanuatu. During the expedition, the scientists KEITH P., WATSON R. E. & MARQUET G. 2002. — operated under the “Forests, Mountains, Rivers” Stenogobius (Insularigobius) yateiensis, a new species theme coordinated by Bruno Corbara, Philippe of freshwater goby from New Caledonia (Teleostei: Keith and Jérôme Munzinger. Gobioidei). Bulletin français de Pêche et de Pisciculture We thank also the following for help and assist- 364: 187-196. KEITH P., MARQUET G. & WATSON R. E. 2004a. — Schis- ance during the various trips and expeditions to matogobius vanuatuensis, a new species of freshwater Vanuatu and New Caledonia: É. Vigneux (ONEMA, goby from Vanuatu (Teleostei: Gobioidei). Cybium France), C. Flouhr (Hytec), G. Ségura (ISE), C. 28 (3): 237-241. Lord (MNHN), F. Busson (MNHN), L. Dehar- KEITH P., WATSON R. E. & MARQUET G. 2004b. — veng (MNHN), P. Gerbeaux (IUCN), J. J. Cassan Sicyopterus aiensis, a new species of freshwater goby (Teleostei: Gobioidei) from Vanuatu, South Pacifi c. (North Province, New Caledonia), the Environment Cybium 28 (2): 111-118. Unit of Vanuatu, especially D. Kalfatak, R. Nari, KEITH P., GALEWSKI T., CATTANEO-BERREBI G., HOAREAU J. White and Ernest Banni for their interest and T. & BERREBI P. 2005a. — Ubiquity of Sicyopterus concerns for the conservation of fl ora and fauna lagocephalus (Teleostei: Gobioidei) and phylogeography native to Vanuatu, Mr Alsen Obed and Mr Glen of the genus Sicyopterus in the Indo-Pacifi c area inferred from mitochondrial cytochrome b gene. Molecular Alo, Fisheries Offi ce in Santo; Mr Kenery Alvea, Phylogenetics and Evolution 37: 721-732. Environment Offi cer, Sanma Province; Ms. Rol- KEITH P., HOAREAU T. & B OSC P. 2005b. — A new species lenas Tavue, Vatthe Offi ce, and all people of New of freshwater goby (Pisces: Teleostei: Gobioidei) from Caledonia and Vanuatu on the sites prospected for Mayotte island (Comoros) and comments about the their cordial reception and their assistance in our genus Cotylopus endemic to Indian Ocean. Journal of Natural History 39 (17): 1395-1405. prospections on the rivers. KEITH P., LORD C. & VIGNEUX É. 2006. — In vivo observations on postlarval development of freshwater gobies and eleotrids from French Polynesia and New REFERENCES Caledonia. Ichthyological Exploration of Freshwaters 17: 187-191. AKIHITO 1986. — Some morphological characters KEITH P., MARQUET G. & WATSON R. E. 2007a. — considered to be important in gobiid phylogeny, in Stiphodon kalfatak, a new species of freshwater goby ICHTHYOLOGICAL SOCIETY OF JAPAN (ed.), Indo-Pacifi c from Vanuatu (Teleostei: Gobioidei: Sicydiinae). fi sh biology. Proceedings of the Second International Cybium 31 (1): 33-37. Conference on Indo-Pacifi c Fishes, Tokyo: 629-639. KEITH P., MARQUET G., WATSON R. E. 2007b. — Akihito BOUCHET P., LE GUYADER H. & PASCAL O. 2008. — futuna, a new species of freshwater goby from the South Des voyages de Cook à l’expédition Santo 2006 : un Pacifi c (Teleostei: Gobioidei: Sicydiinae). Cybium renouveau des explorations naturalistes des îles du 31 (4): 471-476. Pacifi que. Journal de la Société des Océanistes 126-127: KEITH P., HOAREAU T. B., LORD C., AH-YANE O., 167-185. GIMONNEAU G., ROBINET T. & VALADE P. 2008. — BOUCHET P., LE GUYADER H. & PASCAL O. (eds) in Characterisation of post-larval to juvenile stages, press. — The Natural History of Santo. Patrimoines metamorphosis, and recruitment of an amphidromous naturels. goby, Sicyopterus lagocephalus (Pallas 1767) (Teleostei: GERBEAUX P., RICHMOND C. & CHADDERTON L. Gobiidae: Sicydiinae). Marine and Freshwater Research 1998. — Report on a trip to Vanuatu. Department 59 (10): 876-889. of environnment unit, Vanuatu, 12 p. KEITH P., LORD C., MARQUET G. & KALFATAK D. 2009. — KEITH P. 2003. — Review paper: biology and ecology Biodiversity and biogeography of amphidromous fi shes

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from New Caledonia, a comparison with Vanuatu, and S. bustamantei (Teleostei: Gobiidae) and a key in GRANDCOLAS P. (ed.), Zoologica Neocaledonica 7. to west African sicydiines. Th e California Academy Biodiversity studies in New Caledonia. Mémoires du of Sciences Gulf of Guinea Expedition (2201). V. Muséum national d’Histoire naturelle 198: 175-183. Proceedings of California Academy of Science (Ser. 4) LEVITON A. E., GIBBS R. H., HEAL E. & DAWSON C. E. 57 (34): 965-980. 1985. — Standards in herpetology and ichthyology: PLANES S. & GALZIN R. 1997. — New perspectives in part I. Standard symbolic codes for institutional biogeography of coral reef fi sh in the Pacifi c using resource collections in herpetology and ichthyology. phylogeography and population genetics approaches. Copeia 1985: 802-832. Vie et Milieu 47 (4): 375-380. LORD C. & KEITH P. 2006. — Th reatened fi shes of the RADTKE R. L., KINZIE III R. A. & SHAFER D. J. 2001. — world: Protogobius attiti (Watson and Pöllabauer, Temporal and spatial variation in length of larval life 1998) (Galaxiidae). Environmental Biology of Fishes and size at settlement of the Hawaiian amphidromous 77: 101-102. goby Lentipes concolor. Journal of Fish Biology 59 (4): LORD C. & KEITH P. 2008. — Th reatened fi shes of the 928-938. world: Sicyopterus sarazini Weber & De Beaufort (Gobii- WATSON R. E., KEITH P. & M ARQUET G. 2001. — Sicyopus dae). Environmental Biology of Fishes 83: 169-170. (Smilosicyopus) chloe, a new species of freshwater goby MARQUET G., KEITH P. & VIGNEUX É. 2003. — Atlas from New Caledonia (Teleostei: Gobioidei: Sicydiinae). des poissons et crustacés d’eau douce de la Nouvelle- Cybium 25 (1): 41-52. Calédonie. Patrimoines naturels 58: 1-282. WATSON R. E., KEITH P. & M ARQUET G. 2002. — Lentipes MCDOWALL R. M. 2007. — On amphidromy, a distinct kaaea, a new species of freshwater goby from New form of diadromy in aquatic organisms. Fish & Fisheries Caledonia (Teleostei: Gobioidei: Sicydiinae). Bulletin 8: 1-13. français de Pêche et de Pisciculture 364: 173-185. MURPHY C. A. & COWAN J. H. 2007. — Production, WATSON R. E., KEITH P. & M ARQUET G. 2005. — marine larval retention or dispersal, and recruitment Stiphodon sapphirinus, a new species of freshwater goby of amphidromous Hawaiian Gobioids: issues and of New Caledonia (Teleostei: Gobioidei: Sicydiinae). implications. Bishop Museum Bulletin of Cultural and Cybium 29 (4): 339-345. Environmental Studies 3: 63-74. WATSON R. E., KEITH P. & M ARQUET G. 2007. — Akihito PEZOLD F., IWAMOTO T. & H ARRISON I. J. 2006. — vanuatu, a new genus and new species of freshwater Multivariate analysis of sicydiines of São Tomé and goby from the South Pacifi c (Teleostei: Gobioidei: Príncipe with redescription of Sicydium brevifi le Sicydiinae). Cybium 31 (3): 341-349.

Submitted on 26 September 2008; accepted on 21 December 2008.

ZOOSYSTEMA • 2009 • 31 (3) 479 Keith P. et al.

APPENDIX

TABLE 1. — Premaxillary teeth in selected species of Stiphodon Weber, 1895.

Premaxillary teeth 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 S. mele n. sp. 111111 S. rubromaculatus 1221 1 S. kalfatak 111 32 S. sapphirinus 13513 45713212 111 1 S. birdsong 1 2681014837213 1 1 S. hydoreibatus 2 2133111 12 1 1 S. astilbos 111111 S. surrufus 111 121 S. tuivi 2 161127147116674422 11

TABLE 2. — Lateral scale counts in selected species of Stiphodon Weber, 1895.

Lateral scales 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 S. mele n. sp. 21112 S. rubromaculatus 1121 2 S. kalfatak 12221 S. sapphirinus 112245593332111 S. birdsong 1 263758654242 222 111 S. hydoreibatus 1 22122231 1 S. astilbos 21121 S. surrufus 111111 S. tuivi 165386911768473211

TABLE 3. — Transverse back scale counts in selected species of Stiphodon Weber, 1895.

Transverse back scales 3 4 5 6 7 8 9 10 11 12 13 14 15 S. mele n. sp. 11321 S. rubromaculatus 241 S. kalfatak 26 S. sapphirinus 40 1 1 S. birdsong 4 3 31416147 1 S. hydoreibatus 313 S. astilbos 7 S. surrufus 2 221 S. tuivi 7432671

TABLE 4. — Transverse forward scale counts in selected species of Stiphodon Weber, 1895.

Transverse forward scales 0 1 2345678910111213141516171819 S. mele n. sp. 32 11 S. rubromaculatus 2212 S. kalfatak 26 S. sapphirinus 236786532 S. birdsong 2 2627713954411 1 S. hydoreibatus 23334 S. astilbos 222 1 S. surrufus 11 32 S. tuivi 2 8 72920184 1

480 ZOOSYSTEMA • 2009 • 31 (3) A new species of goby (Teleostei, Gobiidae) from Vanuatu and New Caledonia

TABLE 5. — Predorsal scale counts in selected species of Stiphodon Weber, 1895.

Predorsal scales 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 S. mele n. sp. ♂ 5 S. mele n. sp. ♀ 11 S. rubromaculatus ♂ 6 S. rubromaculatus ♀ 1 S. kalfatak ♂ 1211 S. kalfatak ♀ 111 S. sapphirinus ♂ 27 1 S. sapphirinus ♀ 31132111 S. birdsong ♂ 32 S. birdsong ♀ 55 S. hydoreibatus ♂ 51 S. hydoreibatus ♀ 2 3 1 22221 S. astilbos ♂ 1111 S. astilbos ♀ 111 S. surrufus ♂ 3 S. surrufus ♀ 21 S. tuivi ♂ 1067862768451 21 S. tuivi ♀ 1 422255231

TABLE 6. — Predorsal length in selected species of Stiphodon TABLE 7. — Preanal length in selected species of Stiphodon Weber, 1895 expressed to the nearest whole percent of standard Weber, 1895 expressed to the nearest whole percent of standard length. length.

Predorsal length 31 32 33 34 35 36 37 38 39 40 41 42 Preanal length 49 50 51 52 53 54 55 56 57 58 59 61 S. mele n. sp. 2111 1 S. mele n. sp. 1 411 S. rubromaculatus 1222 S. rubromaculatus 1222 S. kalfatak 121121 S. kalfatak 11 1311 S. sapphirinus 4 7 12 10 6 2 S. sapphirinus 3211668221 S. birdsong 2 7 20 23 13 2 S. birdsong 1 3 9 610 91110 7 S. hydoreibatus 13251 S. hydoreibatus 222222 S. astilbos 112 1 2 S. astilbos 1231 S. surrufus 2121 S. surrufus 122 1 S. tuivi 10 12 16 20 15 11 2 S. tuivi 357132410109421

TABLE 8. — Head length in selected species of Stiphodon Weber, 1895 TABLE 9. — Jaw length in selected species of Stiphodon Weber, 1895 expressed to the nearest whole percent of standard length. expressed to the nearest whole percent of standard length.

Head length 18 19 20 21 22 23 24 25 26 Jaw length 6 7 8 9 10 11 S. mele n. sp. 21121 S. mele n. sp. 1231 S. rubromaculatus 34 S. rubromaculatus 124 S. kalfatak 112121 S. kalfatak 1124 S. sapphirinus 11 18 9 3 S. sapphirinus 11 26 4 S. birdsong 52522115 S. birdsong 637231 S. hydoreibatus 1434 S. hydoreibatus 1631 S. astilbos 11113 S. astilbos 214 S. surrufus 24 S. surrufus 1221 S. tuivi 1 1 22 35 27 10 2 S. tuivi 18 37 23 9 1

ZOOSYSTEMA • 2009 • 31 (3) 481 Keith P. et al.

TABLE 10. — Caudal peduncle length in selected species of Stiphodon Weber, 1895 expressed to the nearest whole percent of standard length.

Caudal peduncle length 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 S. mele n. sp. 221 1 1 S. rubromaculatus 2221 S. kalfatak 122111 S. sapphirinus 65159411 S. birdsong 1 5 12 25 15 10 2 S. hydoreibatus 2271 S. astilbos 21 121 S. surrufus 11 4 S. tuivi 3 6 12 28 22 10 5 1

TABLE 11. — Caudal peduncle depth in selected species of Stiphodon TABLE 12. — Body depth at second dorsal origin in males in selected Weber, 1895 expressed to the nearest whole percent of standard species of Stiphodon Weber, 1895 expressed to the nearest whole length. percent of standard length.

Caudal peduncle depth 7 8 9 10 11 12 Body depth at second 910111213141516 S. mele n. sp. 133 dorsal origin in males S. rubromaculatus 115 S. mele n. sp. 13 1 S. kalfatak 242 S. rubromaculatus 132 S. sapphirinus 214205 S. kalfatak 23 S. birdsong 336332 S. sapphirinus 211112 S. hydoreibatus 75 S. birdsong 20 11 1 S. astilbos 52 S. hydoreibatus 11 S. surrufus 141S. astilbos 3 S. tuivi 4175413 S. surrufus 111 S. tuivi 36202274

TABLE 13. — Second dorsal fi n length in selected species of Stiphodon Weber, 1895 expressed to the nearest whole percent of standard length.

Second dorsal fi n length 26 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 S. mele n. sp. ♂ 1112 S. mele n. sp. ♀ 11 S. rubromaculatus ♂ 11 111 1 S. rubromaculatus ♀ 1 S. kalfatak ♂ 311 S. kalfatak ♀ 12 S. sapphirinus ♂ 1 1 32 1733331 S. sapphirinus ♀ 33511 S. birdsong ♂ 1 3586421 1 S. birdsong ♀ 29995 1 S. hydoreibatus ♂ 11 S. hydoreibatus ♀ 14121 S. astilbos ♂ 12 1 S. astilbos ♀ 11 1 S. surrufus ♂ 111 S. surrufus ♀ 11 1 S. tuivi ♂ 1 1 9 8 7 11 13 5 4 5 S. tuivi ♀ 154741

482 ZOOSYSTEMA • 2009 • 31 (3) A new species of goby (Teleostei, Gobiidae) from Vanuatu and New Caledonia

TABLE 14. — Anal fi n length in selected species of Stiphodon Weber, 1895 expressed to the nearest whole percent of standard length.

Anal fi n length 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 S. mele n. sp. ♂ 211 S. mele n. sp. ♀ 11 S. rubromaculatus ♂ 221 1 S. rubromaculatus ♀ 1 S. kalfatak ♂ 13 1 S. kalfatak ♀ 12 S. sapphirinus ♂ 11 1 514553 2 S. sapphirinus ♀ 11333 2 S. birdsong ♂ 168764 S. birdsong ♀ 1 1 412125 1 S. hydoreibatus ♂ 11 S. hydoreibatus ♀ 1 1223 1 S. astilbos ♂ 111 1 S. astilbos ♀ 11 1 S. surrufus ♂ 12 S. surrufus ♀ 11 1 S. tuivi ♂ 4338138137231 S. tuivi ♀ 225222412

TABLE 15. — Caudal fi n length in selected species of Stiphodon Weber, 1895 expressed to the nearest whole percent of standard length.

Caudal fi n length 17 18 19 20 21 22 23 24 25 26 27 28 S. mele n. sp. ♂ 112 S. mele n. sp. ♀ 2 S. rubromaculatus ♂ 11 3 1 S. rubromaculatus ♀ 1 S. kalfatak ♂ 112 1 S. kalfatak ♀ 11 1 S. sapphirinus ♂ 2 2351122 1 S. sapphirinus ♀ 1 25231 S. birdsong ♂ 133222 S. birdsong ♀ 11861 S. hydoreibatus ♂ 11 S. hydoreibatus ♀ 133111 S. astilbos ♂ 31 S. astilbos ♀ 12 S. surrufus ♂ 12 S. surrufus ♀ 12 S. tuivi ♂ 1 1 2 19 13 14 10 4 1 S. tuivi ♀ 26842

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